Brazil nut fractions cause histopathological damage in the midgut of the maize weevil Sitophilus zeamais (Coleoptera: Curculionidae)

Frações de castanha-do-brasil causam danos histopatológicos no intestino médio do gorgulho do milho Sitophilus zeamais (Coleoptera: Curculionidae)

Abstract

Some pests, such as those of the genus Sitophilus, are potentially harmful due to their voracity, high biotic potential, and the fact that they carry out cross-infestation (in this case, insects moving from the field to the warehouses). The objective was to understand the effects caused by Brazil nut fractions on the proventriculus and midgut of the species Sitophilus zeamais, aiming to evaluate whether this Amazonian product can favor the control of this species. Consequently, it was found that in treatments T3 (pig feed + in natura nuts) and T5 (pig feed + defatted nut fraction), where the organic fractions of the Brazil nut were preserved, there were occurrences of damage, thus affecting the structure of the midgut through the observation of cytoplasmic vacuolization, release of cellular fragments into the lumen, and severe morphological changes in the shift from columnar to cubic cell shapes, respectively. These lesions can be considered capable of leading the insect to death, since these compromised midgut cells hinder nutrient absorption by the insect, which makes Brazil nut a natural product that contains potential compounds to be used in the control of S. zeamais.

Keywords:
intestinal cells; foregut; midgut; pest; proventriculus; vacuolization

Resumo

Algumas pragas, como por exemplo as do gênero Sitophilus são potencialmente prejudicais devido a sua voracidade, seu elevado potencial biótico e ao fato de realizarem a infestação cruzada (nesse caso, insetos vindos do campo para os armazéns). O objetivo foi compreender os efeitos causados por frações da castanha-do-brasil no proventrículo e no intestino médio da espécie Sitophilus zeamais visando avaliar se esse produto amazônico pode favorecer o controle dessa espécie. Com isso, foi verificado que, nos tratamentos T3 (ração suína + castanha in natura) e T5 (ração suína + castanha desengordurada), onde foram preservadas as frações orgânicas da castanha-do-brasil, houve ocorrências de danos, afetando assim a estrutura do intestino médio pela constatação de vacuolização citoplasmática, liberação de fragmentos celulares no lúmen e alterações morfológicas severas na mudança do formato de células colunares para cúbicas, respectivamente. Essas lesões podem ser consideradas capazes de levar o inseto a morte, uma vez que essas células do intestino médio comprometidas dificulta a absorção de nutrientes pelo inseto o que faz com que a castanha-do-brasil possa ser considerada um produto natural que apresenta compostos potenciais para serem usados no controle de S. zeamais.

Palavras-chave:
células intestinais; intestino anterior; intestino médio; praga; proventrículo; vacuolização

1. Introduction

Sitophilus zeamais Motschulsky, 1885 (Coleoptera: Curculionidae) is a stored product pest with high biotic potential, damaging grains (Alves and Poltronieri, 2024; Radünz et al., 2024; Ojo and Omoloye, 2012; Antunes et al., 2011; Fragoso et al., 2005; Caneppele et al., 2003; Faroni, 1992). These insects damage cereals in the field (Likhayo and Hodges 2000; Boiça Junior et al., 1996), during transport and storage (Lorini et al., 2015; Gallo et al., 2002).

Grains damaged by insect pests have low nutritional quality (Ojo and Omoloye, 2012), in addition to susceptibility to the development of secondary and opportunistic organisms, such as fungi and bacteria (Marsaro Júnior et al., 2005; Norambuena et al., 2016), compromising the final product, as occurs with feed intended for production animals (Aquino and Potenza, 2013; Stringhini et al., 2000).

Chemical insecticides are the most widely used strategy for the control of S. zeamais; however, they can select resistant insect populations, cause damage to stored products, and leave residues harmful to humans and animals (Carneiro et al., 2025; Haddi et al., 2018; Pinto et al., 2016), which increases the need for alternative methods to control stored grain pests (Greco et al., 2024). In this sense, studies on insect-plant interactions have been important for the development of integrated pest management strategies, including the use of bioactive compounds extracted from plants, which can affect the reproduction, development, and mortality of pest insects (Pinto-Zevallos et al., 2013).

Brazil nut kernels are rich in bioactive compounds, such as phenols, flavonoids, and saponins (Costa and Jorge, 2011), which appear to have an anti-feeding effect on S. zeamais (Socreppa et al., 2025; Pires et al., 2020).

Previous investigation showed that the inorganic elements selenium, barium, and bromine, found in large quantities in Brazil nuts, do not cause mortality in S. zeamais (Socreppa et al., 2025). However, this insect may suffer side effects from ingesting the bioactive compounds and inorganic elements present in Brazil nuts, which needs to be investigated. The digestive system is critical for beetle health because it is the site of contact with pathogens and xenobiotics (Serrão et al., 2022). Curculionidae beetles have the alimentary canal divided into foregut, midgut, and hindgut.

The foregut, of ectodermal origin, is formed by a simple epithelium of flattened cells lined by a thick cuticle, differentiated into pharynx, esophagus, crop, and proventriculus, responsible for ingestion, transport, storage, grinding, and pre-digesting food, with the proventriculus being the most differentiated region among insects (Serrão and Santos, 2026).

The midgut, of endodermal origin, is the main organ for digestion and nutrient absorption (Caccia et al., 2019), with digestive cells being responsible for the production of digestive enzymes and absorption (Terra and Ferreira, 2020).

The hindgut has also an ectodermal origin with a single layered epithelium lined by a permeable cuticular intima, since its main organs, ileum and rectum, play essential role in insect homeostasis absorbing salts and water from the digested food and primary urine (Serrão and Santos, 2026).

Knowledge of the morphology of the insect's foregut and midgut is important for a better understanding of its physiological processes and for the development of pest control strategies. Due to the economically important impact of S. zeamais infestation, the objective was to understand the effects caused by Brazil nut fractions added to the morphology of the proventriculus and midgut of this pest to analyze whether this Amazonian product may contain components with potential to control S. zeamais.

2. Material and Methods

2.1. Insects

Non-sexed adults of S. zeamais were obtained from the mass rearing at the Laboratory of Pests and Vectors of the Amazon Cerrado (LAPVAC) of the Federal University of Mato Grosso, Campus of Sinop (11° 50’ S, 55° 38’ W). The insects were kept under laboratory conditions at 25 ± 2 °C, in glass containers protected with Organza-type fabric and fed corn bran.

2.2. Bioassay

Brazil nut fractions were used in the treatments: in natura almond, almond oil, defatted almond, and mineral residue of the almond. Seeking to follow the same methodology as the study that evaluated the survival of Sitophilus zeamais (Socreppa et al., 2025), these fractions were established as follows: peeled and dried in natura almonds were purchased from retail stores in Sinop, Mato Grosso, and processed in a multiprocessor (Oster® or Ninja®) for fractionation. For the extraction of oil from the in natura almonds, 5 g of crushed almonds were weighed and placed in paper filter cartridges. Then, the cartridges were taken to the oil extractor (MARCONI, MA 044/8/50) with 100 mL of petroleum ether P.A., for a period of four hours after the start of the boiling process. The extracted oily fraction was taken for ether evaporation, in a fume hood, for a period of six hours, and then kept in a forced circulation oven at 55-65 °C for 4 hours to remove residual ether. The defatted almond fraction (cake) that remained retained in the cartridge was kept in an oven at 55-65 °C for 4 hours for drying. The fixed mineral residue was obtained from crushed in natura almonds placed in 50 mL porcelain crucibles, which were first taken to a forced circulation oven at 105 °C for 24 hours to remove all water contained in the material. Then, the crucibles containing the dry samples were taken to a heating plate for combustion of part of the organic fraction. After this step, the material was taken to a muffle furnace with ramp heating, starting from 150 °C, with an increase of 50 °C every 30 minutes until reaching 600 °C, remaining for four hours for total burning of the material, leaving only the ash with the mineral fraction (Silva and Queiroz, 2006).

The mixtures were defined as follows, based on the percentages naturally found in the nut: nuts in natura 1:1 (substrate:nut); nut oil 1:0.6 (60% oil); defatted nut fraction 1:1 and nut mineral fraction 1:0.033 (3.3% ash). Thus, six treatments were established, as follows: T1 (corn bran); T2 (commercial pig feed); T3 (pig feed + in natura nuts); T4 (pig feed + 60% nut oil); T5 (pig feed + defatted nut fraction); T6 (pig feed + fixed mineral residue from nut). Each treatment consisted of three replicates, each using 20 non-sexed adults of S. zeamais. After five days in contact with the substrate, five insects from each treatment were randomly removed for histopathological analyses.

2.3. Histopathology

Adult S. zeamais insects were dissected in 125 mM NaCl and the proventriculus and midgut were transferred to Zamboni's fixative solution for 24h at 4 °C (Stefanini et al., 1967). Subsequently, the samples were dehydrated in an increasing ethanol series (70%, 80%, 90% and 95%) and embedded in Leica historesin following the manufacturer's instructions. Sections 3µm thick were obtained with a Leica RM2245 rotary microtome, stained with buffered Toluidine Blue, and analyzed and photographed under a light microscope (Olympus CX31) coupled with a Nikon D3100 camera.

3. Results

The proventriculus of S. zeamais adults has a wall formed by eight projections (lips) into the lumen, with the surface of the simple flattened epithelium lined by a thick intima with spine-like projections, and well-developed external, longitudinal and circular muscle layers (Figure 1A). No morphological alterations were observed in the proventriculus of insects fed with the Brazil nut fractions (Figure 1B).

Figure 1
(A) and (B) Light micrograph of a cross-section of the proventriculus of Sitophilus zeamais (Coleoptera: Curculionidae), showing flattened epithelium (ep), thick cuticle (ct) and well-developed muscles (m), lumen (L).

The midgut of S. zeamais in the control treatment was formed by a simple epithelium of columnar cells with a well-developed nucleus rich in decondensed chromatin (Figure 2A). Furthermore, the cytoplasm of the cells showed some strongly basophilic regions with a few vacuoles (Figures 22B). In the apical region of the epithelium, the presence of a well-developed brush border was observed (Figures 22B).

Figure 2
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in the control. (A) epithelium with columnar digestive cells (CD) showing apical brush border (Bb), nucleus (N) cytoplasm with basophilic region (arrowhead); (B) Epithelium showing apical brush border (Bb), cytoplasm with vacuoles (Va) and nucleus with clumps of decondensed chromatin (N).

The midgut epithelium of insects from treatment T2 showed morphological alterations with increased abundance of cytoplasmic vacuoles, nuclei with condensed chromatin and some pyknotic ones (Figures 3AC). Some cellular fragments, some containing nuclei, were released into the intestinal lumen (Figures 3BD), but in some regions the striated border was preserved (Figure 3B).

Figure 3
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in treatment T2. (A) Epithelium (ep) with digestive cells (CD); (B) Brush border (Bb), nucleus (N) with condensed chromatin and pyknotic nucleus (Np); (C) and (D) Cells in the lumen cavity (arrowhead), nucleus (N).

In treatment T3, the epithelium showed cytoplasmic vacuolization, but the nuclei did not show significant changes compared to the control insects, being well-developed with decondensed chromatin (Figures 4AD). Cellular fragments were found in the midgut lumen (Figures 4AD) and the brush border showed signs of disorganization in this treatment (Figures 4BD).

Figure 4
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in treatment T3. (A) Epithelium (ep) with digestive cells in the cavity (CD) (arrowhead); (B-D) Cell fragments (arrowhead) in the lumen cavity, cell nucleus (N) with decondensed chromatin, absence of brush border (dotted arrow), cytoplasm with vacuoles (Va).

In treatment T4, the midgut epithelium of S. zeamais was similar to that observed in the control (Figures 5AC), although a few cellular fragments occurred in the lumen (Figures 55B).

Figure 5
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in treatment T4. (A-C) epithelium with columnar digestive cells (CD) showing apical brush border (Bb), nucleus (N) and cellular fragments (arrowhead) in the lumen cavity.

In treatment T5, the epithelium showed severe histopathological damage with a change in shape from columnar to cubic (Figures 6AD), occurrence of large cytoplasmic vacuoles (Figures 6AD); disorganized brush border (Figures 66C) and abundant cellular fragments in the lumen (Figures 66D).

Figure 6
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in treatment T5. (A) epithelium with columnar digestive cells (CD); (B-D) epithelium with digestive cells (CD) with disorganized brush border (dotted arrow), large cytoplasmic vacuoles (Va), cell fragment (Fc).

The midgut epithelium of the beetles in treatment T6 also showed morphological alterations compared to the control, with the occurrence of cytoplasmic vacuoles, cellular fragments in the lumen, and disorganization of the brush border (Figures 7AD).

Figure 7
Light micrographs of the midgut in adults of Sitophilus zeamais (Coleoptera: Curculionidae) in treatment T6. (A-D) epithelium with columnar digestive cells (CD) with absence of brush border (dotted arrow), cytoplasmic vacuoles (Va), cellular fragments (arrowhead) in the lumen, nucleus (N), muscle tissue (m).

4. Discussion

The proventriculus of S. zeamais is similar to that reported for coleopterans, with a thick cuticle presenting spine-like projections and developed musculature, indicating a function in food grinding (Sousa et al., 2013; Rubio et al., 2008; Baker et al., 1984).

The muscle fibers of the foregut of S. zeamais are organized and thick, agreeing with the characterization made for this species kept on its natural food substrate (corn) (Sousa, 2013), and also for other curculionids such as Aegorhinus superciliosus (Coleoptera: Curculionidae) collected in the field in blueberry plantations (Medel et al., 2013), Eusomus ovulum (Coleoptera: Curculionidae) collected on plants of the genus Trifolium (Fabaceae) (Koçakoğlu et al., 2020), Epiphaneus malachiticus (Coleoptera: Curculionidae) collected in the field (Candan et al., 2019) and for Odoiporus longicollis (Coleoptera: Curculionidae) collected in banana plantations (Singh and Prasad, 2013). It is worth noting that the muscles surrounding the foregut and also the hindgut are normally more developed, as they function as dilators (Holtof et al., 2019).

The preservation of the proventriculus structure indicates that the different treatments do not affect this region of the digestive system of S. zeamais adults, which may be due to the function of this intestinal region being food grinding, featuring developed muscles and a thick cuticle with spine-like projections (Serrão and Santos, 2026; Szinwelski et al., 2009), which may ensure the integrity of this part of the intestine.

The columnar epithelial cells with well-developed nuclei with a predominance of decondensed chromatin in the midgut of S. zeamais in the control indicate that transcription processes (mRNA formation) are occurring normally (Vélez Arango et al., 2024; Maori et al., 2019), characteristic of cells that produce and secrete digestive enzymes and absorb nutrients. The cytoplasm of these digestive cells with some basophilic regions indicates functional characteristics for protein synthesis and secretion and absorption (Sousa and Conte, 2013). The presence of some vacuoles in the epithelial cells is similar to that reported in the midgut of this insect fed corn (Sousa and Conte, 2013). On the apical surface of the epithelium, the brush border is evident, important for increasing the cell surface for the release of digestive enzymes and the assimilation of nutrients (Huang et al., 2015).

The morphological alterations observed in the midgut of S. zeamais fed commercial pig feed (treatment T2), which resulted in epithelial cells with cytoplasmic vacuolization, compact and pyknotic nuclei, and release of cellular fragments into the intestinal lumen, suggest the occurrence of autophagy and cell death by apoptosis. Autophagy is involved in the cell renewal process, which is an important process for homeostasis triggered in cases of response to nutrient deprivation or stress, and can be considered a cell survival mechanism (Reis et al., 2024; Madeo et al., 2015; Wirawan et al., 2012). However, in excess, autophagy can trigger cell death by apoptosis (Peng et al., 2025), as evidenced by the occurrence of cells with nuclear pyknosis and release of cellular fragments (Cruz et al., 2010; Taatjes et al., 2008; Hacker, 2000).

The exposure of S. zeamais to commercial pig feed + nuts in natura (treatment T3) promoted cytoplasmic vacuolization and release of cellular fragments into the lumen, indicating a potential detoxification mechanism (Serra et al., 2023; Domingues et al., 2020; Santos-Junior et al., 2020; Carneiro et al., 2019). In bees, alterations in the midgut were observed when they were exposed to fungicides (Serra et al., 2023) and to insecticides (Motta et al., 2023). The disorganization in some regions of the brush border, observed in the midgut that was on the substrate containing feed and in natura Brazil nut, indicates significant morphophysiological alterations, as this structure plays an important role in digestion, nutrient absorption, and ion transport (Nere et al., 2025; Sousa and Conte, 2013). Xenobiotics, such as insecticides, have been reported to cause disorganization of the midgut brush border in insects (Serra et al., 2023; Campos et al., 2021; Santos-Junior et al., 2020), compromising the digestibility and health of these organisms.

Adults of S. zeamais subjected to the treatment of feed containing pig feed + 60% nut oil (T4) did not have evident morphological alterations, which may have been due to a reduction or interruption in food consumption. Even though there were cell fragments in the midgut lumen, this occurred on a small scale and has been reported to occur constitutively in insects. This process can be interpreted as normal physiological events, such as tissue cell renewal, or as a response to the conditions offered to the insects. The cells present in the lumen cavity can be verified, as in adult insects, the intestinal epithelium is one of the few highly regenerative tissues (Zhang and Edgar, 2022). The excess oil present in the substrate offered to the insects can produce adverse effects on them, as Brazil nut oil contains bioactive substances, such as unsaturated fatty acids, including linoleic and oleic acids (Carvalho et al., 2022). In high concentrations, these phytochemicals can affect insects due to their toxicity, repellency, fumigation, and growth inhibition, mainly by influencing feeding and consequent nutrient acquisition (Singh et al., 2021; Rajashekar et al., 2012), a fact that may have occurred in the present research.

When exposing the insects to treatment T5 (pig feed + defatted nut fraction), the epithelial cells of the midgut showed severe morphological alterations such as changes in shape, from columnar to cubic, which can be considered as evidence of damage or stress, since most of the cells that make up the midgut epithelium are columnar (Caccia et al., 2019; Pinheiro et al., 2008). Changes in cell shape and cellular rearrangements result in dynamic changes in the packing of epithelial cells (Lemke and Nelson, 2021). These alterations have been reported in the columnar cells of the midgut of Plodia interpunctella (Hübner, 1813) (Lepidoptera: Pyralidae) (Abdel-Razek et al., 2002), Bombyx mori Linnaeus1 1758 (Lepidoptera: Bombycidae) (Yao et al., 2008), Alabama argillacea (Hübner, 1818) (Lepidoptera: Noctuidae) (Sousa et al., 2010) and Anticarsia gemmatalis Hübner, 1818 (Lepidoptera: Noctuidae) (Castro et al., 2019) when exposed to the bacterium Bacillus thuringiensis Berliner (Bacillaceae) (Bt), which is a bioinsecticide.

Characteristics such as increased cytoplasmic vacuolization and large vacuoles, degenerated microvilli, and fragmentation of cellular content in the midgut lumen were also observed in A. gemmatalis when exposed to B. thuringiensis (Bt) as an alternative to chemical control (Castro et al., 2019). Exposing Tribolium castaneum Herbst 1797 (Coleoptera: Tenebrionidae) insects to B. thuringiensis (Bt) resulted in alterations such as reduction of microvilli, pyknotic nuclei of columnar cells, and cytoplasmic vacuolization, showing that this coleopteran loses integrity in the midgut cells when in contact with this bacterium (Abdel-Razek et al., 2002). It is possible that the compound present in the defatted fraction of the Brazil nut may affect the epithelial cells of the midgut in a way that reaches potential pathways to cause the death of these cells.

The release of cellular fragments into the midgut lumen of insects observed in treatment T6 has been associated as possible mechanisms of cellular detoxification or cell death by apoptosis, a fact generally caused by stress from some agent such as the mineral fraction of the treatment. Similar results were found in the midgut epithelium of several bee species exposed to commercial insecticide formulations, such as thiamethoxam in Partamona helleri Friese, 1900 (Hymenoptera: Meliponini) (Motta et al., 2023), cyflumetofen (Reis et al., 2024), spiromesifen (Serra et al., 2021) and with the fungicide azoxystrobin in Apis mellifera Linnaeus, 1758 (Hymenoptera, Apidae) (Serra et al., 2023). However, when evaluating the mineral fraction of Brazil nut kernels added to ground feed, those treatments where the presence of minerals considered toxic, such as selenium, bromine, and barium, and even others, did not affect the lifespan of S. zeamais (Socreppa et al., 2025).

5. Conclusion

The Brazil nut fractions do not affect the morphology of the proventriculus in S. zeamais adults; on the other hand, the midgut was characterized by morphological differences with the offered treatments.

The fractions established containing Brazil nuts in natura (pig feed + nuts in natura – T3) and Brazil nuts defatted (pig feed + defatted nut fraction - T5) were those that affected the morphological structures of the midgut, which may be associated with the organic fraction of this almond, which may result in mortality of these insects and thus present potential compounds for the control of this pest due to the occurrence of damage to the midgut cells and, consequently, interfering with nutrient absorption and causing cell death.

Acknowledgements

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior- Brasil (CAPES) and also we are grateful to the “Centro para Inovação e Tecnologia em Bioenergia e Sustentabilidade (CIT)” of the Federal University of Mato Grosso – Sinop, MT. To the “Laboratório de Histotécnica (UFV – Rio Paranaíba)” and to the “Departamento de Biologia Geral (DBG)” both at the Federal University of Viçosa.

Data Availability Statement

All data are available to the journal and can be requested from the corresponding author any time.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  17 July 2026
• Date of issue
  2026

History

• Received
  14 Dec 2025
• Accepted
  25 Mar 2026